A new species of tapaculo (Rhinocryptidae: Scytalopus) from ......Volume 132, 2015, pp. 450–466 DOI: 10.1642/AUK-14-166.1 RESEARCH ARTICLE A new species of tapaculo (Rhinocryptidae:

Volume 132, 2015, pp. 450–466DOI: 10.1642/AUK-14-166.1

RESEARCH ARTICLE

A new species of tapaculo (Rhinocryptidae: Scytalopus) from the Serranıade Perija of Colombia and Venezuela

Jorge Enrique Avendano,1a* Andres M. Cuervo,2,3 Juan Pablo Lopez-O.,4 Natalia Gutierrez-Pinto,1,5

Alexander Cortes-Diago,6 and Carlos Daniel Cadena1

1 Laboratorio de Biologıa Evolutiva de Vertebrados, Departamento de Ciencias Biologicas, Universidad de los Andes, Bogota,Colombia

2 Department of Biological Sciences and Museum of Natural Science, Louisiana State University, Baton Rouge, Louisiana, USA3 Department of Ecology and Evolutionary Biology, Tulane University, New Orleans, Louisiana, USA4 Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogota, Colombia5 Department of Biology, University of Miami, Coral Gables, Florida, USA6 Fundacion EcoHabitats, Popayan, Colombiaa Current address: Programa de Biologıa y Museo de Historia Natural, Universidad de los Llanos, Sede Barcelona, Villavicencio,

Colombia* Corresponding author: [email protected]

Submitted July 14, 2014; Accepted December 22, 2014; Published March 11, 2015

ABSTRACTWe describe Scytalopus perijanus (Perija Tapaculo), a new species in the family Rhinocryptidae (suborder Tyranni) found inhumid montane and elfin forests (1,600–3,225 m elevation) in the Serranıa de Perija of Colombia and Venezuela.Although specimens of this taxon have been available in museums since 1941, they were not carefully studied and wereascribed to different taxa of the latebricola and atratus groups. We obtained a modern series of specimens that, coupledwith analysis of vocal and genetic data, clarified this taxonomic puzzle. The new Scytalopus exhibits distinctivemorphological and vocal traits with respect to all other known species and represents a differentiated evolutionarylineage within a clade of northern species, including S. meridanus, S. caracae, and S. latebricola. The new species has notbeen recorded in sympatry with any other Scytalopus, but it may overlap at lower elevations with S. atratus nigricans,although each uses different microhabitats. Ecological niche modeling indicates that the new species currently has arestricted geographic range within the Serranıa de Perija, where large extents of natural habitat have been cleared andfragmented, particularly on the Colombian slopes. Scytalopus perijanus, however, is uncommon to fairly common inforest fragments, tall secondary forest patches, elfin forest, and paramo vegetation at the treeline.

Keywords: Andes, cloud forest, nest, paramo, systematics, tracheophone suboscines, Scytalopodinae,vocalizations

Una nueva especie de tapaculo (Rhinocryptidae: Scytalopus) de la Serranıa de Perija de Colombia yVenezuela

RESUMENDescribimos a Scytalopus perijanus (Tapaculo de Perija), una nueva especie de la familia Rhinocryptidae (subordenTyranni) del bosque montano alto y paramo (1,600 a 3,225 m) de la Serranıa de Perija de Colombia y Venezuela.Aunque han existido especımenes de este taxon en museos desde 1941, estos no habıan sido estudiados con cuidadoy fueron asignados a diferentes taxones de los grupos latebricola y atratus. Recolectamos una serie moderna deespecımenes que, junto con analisis de datos vocales y geneticos, aclararon este acertijo taxonomico. El nuevoScytalopus exhibe rasgos distintivos morfologicos y vocales con respecto a todas las otras especies conocidas yrepresenta un linaje evolutivo distinto en un clado de especies del norte, incluyendo a S. meridanus, S. caracae y S.latebricola. La nueva especie no ha sido registrada en simpatrıa con ningun otro Scytalopus, pero podrıa solaparse amenores elevaciones con S. atratus nigricans, aunque cada una usa microhabitats diferentes. Los modelos de nichoecologico indican que la distribucion potencial de la nueva especie actualmente esta restringida en la Serranıa dePerija, donde se han perdido o fragmentado amplias areas de habitat natural principalmente en las laderascolombianas. Sin embargo, S. perijanus es poco a moderadamente comun en fragmentos de bosque, en los parches debosque secundario alto, en el bosque enano y en la vegetacion de paramo colindante con el bosque.

Palabras clave: Andes, bosque nublado, nido, paramo, Scytalopodinae, sistematica, suboscines traqueofonos,vocalizaciones

Q 2015 American Ornithologists’ Union. ISSN 0004-8038, electronic ISSN 1938-4254Direct all requests to reproduce journal content to the Central Ornithology Publication Office at [email protected]

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INTRODUCTION

The genus Scytalopus (Rhinocryptidae) includes small,

tracheophone subsocines that inhabit primarily the dense

and humid understory of Neotropical montane forests. At

least 28 species-level Scytalopus taxa have been described

or reclassified in the last 20 years (excluding Brazil; Fjeldsa

and Krabbe 1990, Krabbe and Schulenberg 2003, Remsen

et al. 2014), suggesting that even montane regions that

have been explored to some extent may still harbor

undescribed species in this genus. The Serranıa de Perija is

a mountain spur on the Colombian–Venezuelan border

projecting northward from the Eastern Andes (Cordillera

Oriental) of Colombia. This range is an important area of

endemism for birds and other Neotropical montane

organisms (Vuilleumier 1970, Cracraft 1985, Hernandez-

Camacho et al. 1992), but the composition and evolution-

ary history of the Perija montane avifauna have not been

sufficiently studied, particularly on the Colombian (west-

ern) slope (Phelps 1943, Cuervo et al. 2014, Lopez-O. et al.

2014). Two Scytalopus taxa occur in this range (Gines et al.

1953, Lopez-O. et al. 2014): S. atratus nigricans on the

eastern foothills (Phelps and Phelps 1953) and an

unnamed population (see Lentino et al. 2004) at higherelevations, which we formally describe herein.

In 1941 and 1942, Melbourne A. Carriker, Jr., explored the

western slope of the Serranıa de Perija, collecting 27

specimens of a single species of Scytalopus from 6 localities,

all sent to the Smithsonian Institution National Museum ofNatural History (USNM; C. Ludwig personal communica-

tion). Carriker’s serieswasmistakenly assumed to correspond

to S. [femoralis] atratus nigricans (Carriker 1954, Meyer de

Schauensee 1959), a taxon described from 2 specimens

collected at lower elevations on the Venezuelan side of the

range (Phelps and Phelps 1953). Carriker’s specimens lack the

diagnostic traits of S. atratus nigricans, including size, color

patterns, and a white patch on the middle crown (Phelps and

Phelps 1953, Hilty 2003). From1951 to 1978, a larger series of

the same taxon collected by Carriker was obtained on the

eastern slope by personnel of the Adolfo Pons (PONS) and

Phelps (COP) collections in Venezuela (Gines and Yepez

1953, Aveledo Hostos and Perez Chinchilla 1989).

The combined series of this upper montane Scytalopus

remained unstudied for decades despite its existence in

different museums since 1941. As a result, this tapaculo has

not only been ascribed inconsistently to different taxa over

the years, but it has also been overlooked in the literature on

avian distributions and monographs on tapaculos (e.g.,

Phelps and Phelps 1950, Peters 1951, Meyer de Schauensee

1964, Fjeldsa and Krabbe 1990, Krabbe and Schulenberg

2003, Clements et al. 2014). It was only in 1953 that this

population was explicitly referred to as a member of the

latebricola group (sensu Zimmer 1939); based on specimens

at the PONS collection, it was then identified as S. caracae

(Gines et al. 1953, Gines and Yepez 1953); however, most

later authors referring to this population have treated it

either as meridanus (Phelps and Phelps 1963, Viloria and

Calchi La C 1993, Hilty 2003) or simply as S. latebricola

(Ridgely and Tudor 1994). More recently, it has been

hypothesized that the Perija population could represent

either an undescribed species based on vocal differences

with respect to better-knownVenezuelan taxa (Lentino et al.

2004; C. Sharpe andD.Ascanio personal communication) or

a subspecies related to S. griseicollis or S. meridanus based

on morphology (Donegan and Avendano 2008).

In September 2006, J. P. Lopez-O. and A. Cortes-Diago

collected 2 Scytalopus in cloud forest at 2,450 m on the

western (Colombian) slope of the Serranıa de Perija. No

taxonomic identification was made from the specimens at

the time due to the lack of sufficent comparative material,

tape recordings, or genetic data. Between July 2008 and

February 2009, however, Lopez-O. et al. (2014) collected a

modern series of 16 specimens and sound recordings of

this Scytalopus in cloud forest, elfin forest, and paramo

(2,450 and 3,050 m) in the same general region and near

the area first visited by Carriker. Based on this new

material, we conducted molecular phylogenetic analyses,

which confirmed that the Perija Scytalopus represents a

unique evolutionary lineage highly divergent from all other

Scytalopus species. Vocal, morphological, and ecological

analyses further confirmed that this population represents

a distinct, undescribed species, which we propose to name:

Scytalopus perijanus, sp. nov.

Perija Tapaculo

Tapaculo de Perija

HolotypeThe holotype is an adult male specimen deposited in the

ornithological collection of the Instituto de Ciencias

Naturales at Universidad Nacional de Colombia (ICN),

number 36745; collected from above vereda El Cinco,

Municipality of Manaure, Department of Cesar, on the

western slope of the Serranıa de Perija, Colombia

(10821 050 00N, 72856 051 00W; ~2,450 m elevation). It was

lured by a playback of a recording of its own primary song

and collected on July 10, 2008, by J. E. Avendano (field

number 695). The holotype was found in a tangled edge

of humid montane forest intermixed with Chusquea

bamboo. Tissue samples of the holotype are preserved in

the genetic resources collections of the Museo de Historia

Natural de la Universidad de los Andes (ANDES-BT 760)

and Instituto Alexander von Humboldt (IAvH-BT 11317),

Colombia. Sound recordings of the holotype’s vocaliza-

tions are deposited in the Coleccion de Sonidos Animales,

Instituto Alexander von Humboldt (IAvH-CSA 2801-16).

The Auk: Ornithological Advances 132:450–466, Q 2015 American Ornithologists’ Union

J. E. Avendano, A. M. Cuervo, J. P. Lopez-O, et al. New species of Scytalopus from Colombia and Venezuela 451

Diagnosis: Plumage Coloration and MorphologyThe new species exhibits all the characteristics of the

genus Scytalopus (Ridgway 1911, Krabbe and Schulenberg

1997, Cuervo et al. 2005, Maurıcio et al. 2008). In plumage

coloration, the new species is most similar to the

population of S. griseicollis of the middle sector of the

Eastern Andes, but adult males and females exhibit a

brown nuchal patch that contrasts with a gray back. Also,

all sexes and ages exhibit a variable extent of buff feathers

in the lower belly. In comparison to the northernmost

populations of S. griseicollis (heretofore referred to as

northern S. griseicollis) from the Tama massif to the

western slope in northern Santander (Zimmer 1939,

Donegan and Avendano 2008), the new species is duller

gray, and its upperparts are not extensively brown.

Scytalopus perijanus is also similar to S. caracae and S.

meridanus but is duller ventrally than S. caracae and has a

gray back, not brown as in S. meridanus. Also, males of the

new species tend to be larger in body and bill size than S.

meridanus (Table 1). Compared to S. latebricola, S.

perijanus can be distinguished by its decidedly brighter

plumage, smaller size, shorter tarsus, and smaller bill (i.e.

shorter in length and height). Compared to S. atratus and

S. sanctaemartae, the new species lacks a white coronal

patch, its dorsal plumage is lighter grayish brown instead

of blackish gray or black, it has no whitish tips to any of the

ventral feathers, and it has a darker throat. The new species

is distinguishable from S. spillmanni and S. parkeri by its

lighter plumage and smaller size (Table 1), and it is lighter

in plumage coloration than S. latrans, with a buffy belly

and extensive brown in the rump and flanks. Nestlings and

fledglings of S. perijanus appear to be distinguishable from

those of northern S. griseicollis by their mostly yellowish

instead of whitish appearance and by a blacker barring; in

S. meridanus, barring in juveniles is less yellowish and

darker brown than in S. perijanus.

Diagnosis: VocalizationsScytalopus perijanus is diagnosable by its vocalizations from

other Scytalopus taxa in multiple temporal and spectral

traits. The primary song of S. perijanus is a short ‘‘churr’’delivered in a series of 2 and up to 65 repeats at 0.5–3.0 s

intervals (Figure 1). Each song churr (0.8 6 0.2 s, 11.9 6 1.5

notes) is distinguishable from the much longer and richer

songs of S. meridanus (20.9 6 7.5 s, 169.8 6 38.6 notes), S.

latebricola (9.9 6 2.3 s, 117.0 6 45.4 notes), and northern

S. griseicollis (1.5 6 0.2 s, 34.6 6 5.5 notes). The song pace

(notes s�1) in S. perijanus (14.7 6 2.1; Figure 2A) is similar

to that of S. meridanus (9.5 6 5.0; Figure 2C), but it is faster

than in S. latebricola (11.5 6 2.3; Mann-Whitney test: U¼8, p , 0.05; Figure 2B) and slower than northern S.

griseicollis (23.7 6 0.9; U ¼ 0, p , 0.01; Figure 2D).

Compared to S. caracae and S. atratus, the new species

songs are distinct in multiple characteristics (Figures 2E,

2F). On spectral traits, S. perijanus can be distinguished

from northern S. griseicollis by a higher minimum frequency

(2.7 6 0.3 vs. 1.7 6 0.5 kHz), maximum frequency (3.8 6

0.2 vs. 2.1 6 0.5 kHz), peak frequency (3.4 6 0.2 vs. 1.9 6

0.5 kHz), and bandwith (frequency range; 1.1 6 0.2 vs. 0.4

6 0.1 kHz; U ¼ 0, p , 0.01), and from S. latebricola by a

lower maximum frequency (3.8 6 0.2 vs. 4.2 6 0.2 kHz; U

¼ 4, p , 0.01). Apparent differences in maximum frequency

and bandwith with respect to S. meridanus (see Discussion)

were not significant based on our sample (U¼ 19, p¼ 0.56);

however, S. perijanus lacks the high-pitched and introduc-

tory notes often included in the songs of both S. meridanus

TABLE 1. Body mass (g) and morphometric measurements (mm) of males of S. perijanus and selected Scytalopus species. Values arepresented as mean and range (in parenthesis). Measurements of S. perijanus are from Colombia and Venezuela; those of S. griseicollisare from the northern sector of the Eastern Andes in Santander and Norte de Santander, Colombia. Data for S. spillmanni andS. parkeri are from Ecuador and taken from Krabbe et al. (2005). See Appendix A for a list of specimens examined.

Species Body mass Wing (flat) Tail Tarsus Bill lengtha Bill heightb Bill widthb

S. perijanus 17.7 (16.5–20.5)n ¼ 10

57.4 (51.5–65.0)n ¼ 32

40.3 (32.0–48.0)n ¼ 32

21.1 (19.4–22.4)n ¼ 32

6.8 (5.7–8.3)n ¼ 32

3.5 (3.0–3.9)n ¼ 27

2.9 (2.1–3.4)n ¼ 32

S. latebricola 23.8 (21.5–26.0)n ¼ 3

62.5 (59.5–66.5)n ¼ 14

42.3 (39.1–45.3)n ¼ 12

23.4 (22.4–24.4)n ¼ 14

7.8 (6.9–8.9)n ¼ 13

4.1 (3.7–4.7)n ¼ 11

3.2 (2.6–3.7)n ¼ 14

S. meridanus 15.3 (13.5–16.5)n ¼ 6

54.0 (50.0–57.0)n ¼ 13

39.9 (35.0–46.0)n ¼ 13

21.2 (19.3–22.6)n ¼ 13

6.0 (5.5–6.6)n ¼ 13

3.1 (2.8–3.5)n ¼ 12

2.5 (2.1–2.6)n ¼ 12

S. caracae — 55.9 (53.0–60.0)n ¼ 16

40.4 (37.0–47.0)n ¼ 16

22.0 (20.8–23.1)n ¼ 16

7.0 (6.5–7.4)n ¼ 16

3.5 (3.2–3.8)n ¼ 16

2.8 (2.5–3.0)n ¼ 16

S. griseicollis 16.6 (15.0–18.5)n ¼ 15

58.0 (52.0–61.0)n ¼ 17

41.4 (36.0–44.7)n ¼ 15

21.2 (19.5–22.2)n ¼ 17

7.1 (6.3–7.9)n ¼ 16

3.3 (3.0–3.7)n ¼ 14

3.2 (2.8–3.6)n ¼ 17

S. spillmanni 25.2 (21.0–30.0)n ¼ 36

61.9 (56.0–67.0)n ¼ 38

45.3 (39.0–54.0)n ¼ 35

24.5 (22.2–26.0)n ¼ 18

7.0 (6.1–7.6)n ¼ 14

— —

S. parkeri 22.5 (21.0–24.4)n ¼ 13

62.9 (59.0–66.0)n ¼ 14

44.5 (42.3–50.0)n ¼ 13

24.6 (23.8–25.3)n ¼ 14

6.4 (6.1–7.0)n ¼ 8

— —

a From fore edge of operculum to tipb At fore edge of operculum


452 New species of Scytalopus from Colombia and Venezuela J. E. Avendano, A. M. Cuervo, J. P. Lopez-O, et al.

and S. latebricola. In addition, the song of S. perijanus is

mainly formed by upstroke notes (as in S. latebricola),

unlike the mostly downstroke or short unmodulated notes

of S. meridanus, and the up-down strokes of S. caracae and

northern S. griseicollis (Figure 2).

The call of S. perijanus is short (0.8 6 0.1 s) and fast-

paced (23.7 6 1.4 notes s�1; Figure 3A); the pace is faster

than in S. latebricola (20.7 6 1.6 notes s�1; U ¼ 23, p ,

0.001; Figure 3B) and S. meridanus (18.3 6 1.5 notes s�1;

U¼ 0, p , 0.001; Figure 3C), but slower than in northern

S. griseicollis (29.3 6 1.9 notes s�1; U¼ 0, p , 0.01; Figure

3D). It is also shorter than in S. meridanus (1.3 6 0.2 s; U

¼ 0, P , 0.001) and has fewer notes than in northern S.

griseicollis (18.9 6 1.6 vs. 27.2 6 5.7; U¼ 9.5, p , 0.001).

Additionally, the call of S. perijanus can be distinguished

from calls of S. latebricola and northern S. griseicollis by

having lower and higher frequency traits, respectively, as

follows: maximum frequency (4.6 6 0.2 vs. 6.2 6 0.5 and

3.2 6 0.2 kHz), minimum frequency (3.4 6 0.1 vs. 4.5 6

0.4 and 2.5 6 0.2 kHz), bandwith (1.3 6 0.1 vs. 1.7 6 0.5

and 0.7 6 0.1 kHz), and peak frequency (4.1 6 0.2 vs. 5.2

6 0.5 and 2.9 6 0.2 kHz; U ¼ 0, p , 0.001 for all

comparisons; Figure 3B–3D). However, the calls of S.

meridanus and of the new species seem not to differ in

spectral traits. The call of S. perijanus is clearly distin-

guishable from those of S. caracae, S. spillmanni, S.

parkeri, and S. atratus in numerous vocal traits (Figure

3E–3H). Likewise, alternative and advertising songs are

distinctive from those known for other Scytalopus (see

Discussion). Distinctiveness of vocal signals across Scyta-

lopus matches phylogenetic clusters (Arctander and

Fjeldsa 1994, Cuervo et al. 2005, Maurıcio et al. 2014;

see Discussion: Systematics); therefore, the new species is

diagnosable morphologically, vocally, and also genetically.

Description of HolotypeThe holotype is a fairly small tapaculo (18.5 g) with 8

rectrices. Lores, forehead, crown, auriculars, mantle, and

scapular area are Dark Neutral Gray 83 (color nomencla-

ture and numbers follow Smithe 1975, 1981). Feathers in

FIGURE 1. Waveform (above) and spectogram (below) depicting the relative amplitude and frequency, respectively, of an excerpt of9 churr phrases of the primary song of S. perijanus sp. nov. recorded at San Antonio, above Manaure, Cesar, Colombia, February 2009(CSA 2804).



the scapular area have inner webs tinged between Dark

Neutral Gray 83 and Verona Brown 223B whereas outer

webs are Dark Neutral Gray 83. The color of the nape is

between Verona Brown 223B and Dark Drab 119B. Back

and rump fall between Verona Brown 223B and Amber 36,

with some feathers of the back Dark Neutral Gray 83.

Upper-tail coverts are Mars Brown 223A and, like the

rump, barred with Sepia 219. Chin, throat, breast, and

FIGURE 2. Sonograms of fragments of the primary song of Scytalopus perijanus sp. nov. and other northern species in the genus. (A)Three bouts by S. perijanus, type locality, February 10, 2009 (CSA 2803); (B) S. latebricola, Santa Marta, Magdalena, Colombia,February 12, 2007 (Krabbe 2008); (C) S. meridanus, 10 km SE La Azulita, Merida, Venezuela, February 18, 1985 (XC 6236, C. Parrish); (D)reeling song of northern S. griseicollis, California, Santander, Colombia, June 16, 2009 (J. E. Avendano); (E) S. caracae, Colonia Tovar,Aragua, Venezuela, October 11, 2005 (XC 3800, N. Athanas); (F) S. atratus cf. nigricans, Tama N. P., Norte de Santander, Colombia,August 1999 (Alvarez et al. 2007).



center of belly are between Medium Neutral Gray 84 and

Light Neutral Gray 85. Sides of breast are Dark Neutral

Gray 83, whereas the lower breast and center of belly are

washed with whitish (close to Pale Neutral Gray 86). Lower

belly and flanks are between Clay 123B and Tawny 38.

Thighs and under-tail coverts are between Verona Brown

223B and Amber 36, and indistinctly barred blackish.

Rectrices and remiges are Vandyke Brown 121. Margin of

tertials and the tip of the outer web are Mikado Brown

121C. Under-wing coverts are Clay 123B. The tip of the 3

external rectrices has a terminal band between Verona

Brown 223B and Amber 36 and a subterminal band of

Sepia 219. Soft parts in life: irides dark brown; bill dull

horn with maxillary tip and mandibular tomia paler; tarsi

brown with frontal side and toes whitish brown; claws

whitish to grayish bone, hind claw gray; and foot soles pale

yellow. The holotype had abundant subcutaneous fat and

light molt in crown, mantle, and throat, and the sixth right

primary was sheathed. Stomach contained insect remains.

Testes were enlarged (left testis: 7.2 3 4.3 mm; right testis

5.4 3 3.2 mm). Measurements (in mm): bill (from tip to

fore edge of operculum) length: 6.8; bill height: 3.4; bill

width: 3.2; tarsus: 20.3; tail: 44.6; wing (flattened) 61.2 mm.

ParatypesWe designate as paratypes a series of 16 round-skin

specimens at ICN and IAvH-A collected in 2008 and 2009.

Eleven were taken at the type locality between~2,450 m and

2,600m: adult males: ICN 36732, 36798, 36766; adult female:

ICN 36729; subadultmales: ICN 36799, 37069, 37091, 37101;

fledgling males: ICN 36734, 36800; fledgling female: ICN

36801. The remaining 4 paratypes were collected upslope in

paramo vegetation at Sabana Rubia, a few kilometers from

theVenezuela border, Municipality of Manaure, Department

of Cesar, Colombia (10822001 00N, 72853051 00W, ~3,025 m):

adult males: ICN 36838, IAvH-A 15290; adult female: ICN

36857; fledgling female: ICN 36852.

EtymologyThe Latin, English, and Spanish names refer to the

Serranıa de Perija of Venezuela and Colombia, the

mountain range to which the new bird species is endemic.

DISCUSSION

Variation Within the Type SeriesPhenotypic variation in the type series is associated with

age and sex and mainly involves the degree of dullness in

the underparts, the extent of buffy coloration in the belly,

and the presence and extension of a brown nuchal patch.

Adult males are similar ventrally to the holotype, although

2 are duller (ICN 36766 and 36798) and another one is

paler (i.e. light gray, ICN 36732). Only one male specimen

(ICN 36798) exhibits a nuchal patch resembling that of the

holotype. Two specimens (ICN 36838 and 36839) have a

gray nuchal patch tinged with Verona Brown, which is

hardly visible in 3 males (ICN 36732, 36766, and 36799).

Color variation in the flanks and under-tail coverts is

subtle, but 3 adult males (ICN 36760, 36798, and 36838)

are slightly duller than the holotype. Coloration of the

lower belly appears to be invariant across male specimens.

In females, the nuchal patch is more conspicuous than in

adult males, between Verona Brown 223B and Raw Sienna

136 (Figure 4). The underparts resemble that of the type and

other adult male specimens but have a lighter lower-breast

mixed with Pale Pinkish Buff 121D reaching the lower belly

(e.g., ICN 36729) or the upper belly (e.g., ICN 36857).

Subadult males have a light gray washing in the belly, tinged

FIGURE 3. Sonograms of calls of Scytalopus perijanus sp. nov.and other northern species in the genus. (A) S. perijanus, SabanaRubia, Manaure, Cesar, Colombia, July 6, 2008 (CSA 2810); (B) S.latebricola, San Lorenzo, Santa Marta, Magdalena, Colombia,February 12, 2007 (Krabbe 2008); (C) S. meridanus, LomaRedonda-La Aguada, Sierra Nevada N. P., Merida, Venezuela,February 18, 2000 (XC 50486, B. Lopez-Lanus); (D) northern S.griseicollis, California, Santander, Colombia, June 6, 2009 (XC86713, J. E. Avendano); (E) S. caracae, Cortada de Maya road,Colonia Tovar, Aragua, Venezuela, October 11, 2005 (XC 3801, N.Athanas); (F) S. spillmanni, Finca Andalucıa, San Vicente delCaguan, Caqueta, Colombia (Alvarez et al. 2007); (G) S. parkeri,Acanama, Loja, Ecuador, February 13, 1991 (XC 32893, N.Krabbe); (H) S. atratus confusus, Bosque Las Animas, Amalfi,Antioquia, Colombia (Alvarez et al. 2007).



with Pale Neutral Gray 86 (ICN 37091) or with Clay color

123B (ICN 37069). The nuchal patch is barely evident in 2

subadult birds (ICN 37091 and 37101) but is conspicuous in

another (ICN 37069), in which the nuchal patch extends to

the upper back, presumably a retention of juvenile plumage.

Like in the holotype, the back of adult males, females, and

one subadultmale (ICN37069) is graymixedwith brown, but

in 2 subadult males (ICN 37101 and 37091) the back is

uniform gray. Rump coloration is slightly tawnier in one

female (ICN 36857) and in subadult males. The extent of

brown in secondaries is also variable, in some cases restricted

to feather tips as in one adult male (ICN 36732) and

immature males, or covering half of each feather (e.g., adult

male ICN 36838) or all the feather (female ICN 36857).

Regarding fledgling plumage, the male (ICN 36734) has

more densely barred upperparts between Verona Brown

223B andMikado Brown 121C than the female (ICN 36852),

which lacked any barring in the head and back; this suggests

that the female fledgling specimen was older than the male

fledgling. In general, light to moderate body molt and light

symmetric molt in wings and tail feathers were present in

adult specimens, and one subadultmale (ICN37069) still had

first-year secondary feathers. The number of rectrices is

variable in the type series (range 8–12), but 5 of 7 adult males

have 12.

Additional Specimens ExaminedWe examined 140 specimens of 5 Scytalopus taxa,

including 45 additional specimens we identified as S.

perijanus (Appendix A). The historical specimens of S.

perijanus, including Carriker’s series, exhibit variation

within the range observed in the type series.

SystematicsWe used DNA sequence data from 3 S. perijanus

specimens (ICN 36729, 36734, and 36766) to provide a

hypothesis of the systematic affinities of the new species

with respect to the other Scytalopus with which it has been

confused in the literature, namely S. caracae, S. latebricola,

S. griseicollis, S. meridanus, and S. atratus. In addition, we

assessed genetic divergence among these species and

others occurring in northern Colombia and Venezuela.

Sequences of the ND2 mitochondrial gene were generated

for an ongoing diversification analysis of the genus with

nearly complete taxon sampling and dense geographic

coverage across the Northern Andes (C. D. Cadena et al.

personal communication). We used sequences of 3

scytalopodinae tapaculos as outgroups: S. novacapitalis,

Myornis senilis, and Eleoscytalopus indigoticus (Maurıcio

et al. 2008). We truncated the original alignment in

reference to an 843-bp sequence of S. caracae obtainedfrom a toe pad sample (COP-IC1209). New sequences

were deposited in GenBank (accession nos. KM668104-

KM668115); specimens used in the analysis are highlighted

in Appendix A.

Ongoing phylogenetic studies of Scytalopus indicate that

S. perijanus belongs in a clade including several species

from the Northern Andes largely corresponding to the

latebricola group (sensu Zimmer 1939) plus a few recently

described species (C. D. Cadena et al. personal commu-

nication). All the taxa considered here, with the exception

of S. atratus and outgroups, also belong to this clade. We

estimated the ND2 gene tree for our study taxa

implementing a Bayesian analysis in MRBAYES, v3.1.2

(Huelsenbeck and Ronquist 2001). Four independent

analyses consisting of 4 MCMC chains were run for 10

million steps, of which 25% were discarded as burn-in. The

best-fit substitution model (GTRþI) according to the

Akaike’s information criterion calculated using MRMO-

DELTEST v2.3 (Nylander 2004) was implemented in the

analysis. In addition, we conducted a maximum-likelihood

(ML) analysis in RAxML, v7.2.6 (Stamatakis 2006), which

implemented the GTRþC model.

Our analyses indicated that the closest relatives of S.

perijanus are S. meridanus, S. caracae, and S. latebricola

(Figure 4), but they were unable to determine its sister

species, possibly as a result of rapid diversification. The

new species and the 3 species mentioned above form a

clade sister to S. griseicollis (Figure 5). Uncorrected genetic

distances between S. perijanus and its 3 closest relatives

FIGURE 4. Female (left) and male (right) Perija TapaculoScytalopus perijanus sp. nov. attending a nestling (center). Notethe more rufous nuchal patch in the female. Watercolor by JonFjeldsa.



ranged from 8.2% to 9.3%, which exceeds divergence

between some closely related species of Andean tapaculos

(Cuervo et al. 2005, Krabbe and Cadena 2010) and is

similar to the divergence among species in the speluncae

group (Mata et al. 2009, Maurıcio et al. 2014). These levels

of divergence suggest a long period of evolutionary

isolation, which is not unusual in lineages of cloud forest

birds endemic to the Serranıa de Perija and adjacent ranges

(Cadena and Cuervo 2010, Derryberry et al. 2011, Benham

et al. 2015, Valderrama et al. 2014).

VocalizationsWe analyzed 9 vocal variables of primary songs and calls to

assess vocal divergence of S. perijanus and 3 geographically

adjacent Scytalopus taxa, including 2 of its closest relatives

(S. latebricola, S. meridanus, and northern S. griseicollis).

Songs of the more geographically isolated S. caracae were

not analyzed, but previous descriptions (Fjeldsa and

Krabbe 1990, Krabbe and Sculenberg 1997) demonstrated

that its vocalizations differ drastically from those of any

other tapaculo. For each vocalization, we measured ~60%

of notes, sampling equal proportions (20%) at the

beginning, middle, and end of the vocalization, using the

program Raven Pro v1.3 (Charif et al. 2008) in blackman

window type, with a resolution of 512 bands and overlap of

99%. We analyzed the following variables for each

vocalization (single calls and song churrs): number of

notes, song length, note pace, note length, interval length

between notes, minimum frequency, maximum frequency,

bandwidth, and peak frequency. We analyzed 4–9 different

individuals per species and measured and averaged 1–4

vocalizations per individual (Appendix B). We determined

whether taxa were vocally distinguishable in multivariate

space using discriminant analyses based on 8 quantitative

variables (i.e. excluding number of notes). Alternative and

advertising songs (Krabbe and Schulenberg 1997) were

qualitatively compared and described.

Discriminant analysis based on average values demon-

strated that S. perijanus, S. latebricola, S. meridanus, and

northern S. griseicollis are distinguishable in multivariate

space in primary songs (Wilks’s k¼ 0.002, F¼ 112.9, p ,

0.001, n¼ 24 individuals) and calls (Wilks’s k¼ 0.003, F¼

FIGURE 5. Phylogenetic position of S. perijanus sp. nov. within a clade of tropical montane Scytalopus from northern South Americabased on ND2 sequences. The phylogeny is a 50% majority-rule consensus tree from the Bayesian analysis. Numbers at nodesrepresent Bayesian posterior probabilities (below) and maximum-likelihood bootstrap support values (above). Numbers in front ofspecies names are the number of individuals sequenced per species (Appendix A). Genetic distances between species (uncorrected-p) are shown as percentages.



145.8, p , 0.001, n¼31 individuals). All vocalizations were

assigned to their corresponding species (Figure 6). Primary

songs were best discriminated by the canonical function 2

(CF 2), which was associated with all variables except note

length. Bandwidth was the only uninformative variable

across analyses. Calls were discriminated by frequency

variables along CF 1 and by pace and song length along CF

2 (Table 2).

The alternative song of S. perijanus (Figure 7A) was

analyzed based on recordings of 5 different individuals.

This song type is elicited as a response to playback or

during agonistic encounters, presumably by males, while

perching 1.0–1.5 m above ground. It consists of a series of

11–22 accelerating churrs, mostly delivered every 0.1–

0.9 s. The first 2–3 churrs of each song consist of 4–5

notes where the first note is flat and the rest are upstrokes.

Then, gradually, the churrs reach 9–14 up-down strokes,

rarely up to 90 notes. This vocalization may be emitted in

ascending or descending pitch, reaching a peak at 3.7–4.5

kHz. The alternative song of S. latebricola has similar pitch

to that of S. perijanus (Figure 7B) but consists of fewer (2–

7) churrs. The first churr in S. latebricola has 7–13 notes;

the following churrs seem to have more notes on average,

and these are up-strokes. In songs composed of 2 churrs,

the last one may extend to up 70 notes in S. latebricola. To

our knowledge, no recordings of alternative songs are

available for S. griseicollis, S. meridanus, or S. caracae.

An advertising song similar to that of females of several

other species of Scytalopus, including S. spillmanni, is given

by S. perijanus, presumably by females (Figure 7C, 7D).

Our single recording was a series (~9.5 s long) of 32

explosive up-down strokes. The first 4 notes were delivered

in pairs and were high-pitched (4.7–5.7 kHz). From the

fifth note on, notes were single, and the pitch gradually

descended to 5.1 kHz at the 24th note, increasing again at

the end to 5.4 kHz. Time intervals between notes decreased

gradually from 0.63 s (between the first and second note) to

0.08–0.16 s. The female advertising song of S. spillmanni is

apparently lower-pitched on average, slightly slower (3.0 vs.

3.4 notes s�1), and composed only of single notes.

DistributionScytalopus perijanus has been recorded at 19 localities on

both slopes of the Serranıa de Perija (Appendix C). On

the Venezuelan slope, it is known from 9 localities,

ranging in elevation from 1,800 (at Cerro Tetarı, La Teta

or Las Tetas and Las Antenas, Rıo Negro, Zulia; Lentino

FIGURE 6. Multivariate space of vocal signals of S. perijanus sp. nov. and other 3 related species from the first 2 functions ofdiscriminant analysis based on 8 quantitative acoustic variables taken on primary songs (A) and calls (B). Values for primary songsand calls are from different individuals; each one represents one vocalization or the average of 2–4 vocalizations per individual(Appendix B).

TABLE 2. Results of discriminant analysis based on 7 acousticmeasurements of primary songs and calls of 4 Scytalopus speciesfrom the Northern Andes: S. perijanus sp. nov., S. latebricola, S.meridanus, and S. griseicollis. The acoustic variables thatcontributed significantly to the first 2 canonical functions (CF)are highlighted in bold. The cumulative variance explained bythe first 2 CF was 92.7% and 92.2% for songs and calls,respectively. Delta frequency is not included because thisvariable did not contribute to the canonical functions.

Primary songs Calls

Acoustic variable CF 1 CF 2 CF 1 CF 2Song length 0.284 �0.618 �0.063 0.380Pace 0.078 0.543 �0.513 �0.536Note length �0.141 �0.239 0.356 0.284Interval length between

notes 0.143 �0.343 0.199 0.361Low frequency �0.190 �0.375 0.644 �0.134High frequency �0.292 �0.639 0.815 �0.208Peak frequency �0.303 �0.611 0.658 �0.162Variance explained 60.80% 31.90% 75.60% 16.60%



et al. 2004) to 3,120 m (at Cerro Tetarı). On the

Colombian slope, S. perijanus is known from 10 localities

in departments of La Guajira and Cesar ranging from

1,600 m (below San Antonio, above Manaure, Cesar) to

3,225 m (Cerro Pintado, La Guajira). However, the lower

elevational limit remains to be established owing to the

scarcity of fieldwork and to the extensive deforestation

below 1,600–2,000 m.

Scytalopus tapaculos often exhibit elevational replace-

ments with habitat seggregation where their ranges meet.

It is likely that only 2 species occur in Serranıa de Perija, as

in the adjacent Sierra Nevada de Santa Marta where an

atratus lineage (i.e. S. sanctaemartae) occurs in the

foothills and mid elevations and latebricola occurs in the

highlands. Scytalopus perijanus has not been recorded in

sympatry with any other Scytalopus species. It is possible,

however, that its range overlaps at mid elevations (1,500–

1,900 m) with that of S. atratus nigricans, which occupies a

different microhabitat, and it is only known in the Serranıa

de Perija from the east slope (contra Carriker 1954, Meyer

de Schauensee 1959) on the basis of 2 specimens taken at

1900 m (Phelps and Phelps 1953, Hilty 2003). However, S.

perijanus might co-occur with the northern populations of

S. griseicollis (Donegan and Avendano 2008) at the low-

elevation range of Serranıa de los Motilones, which may

harbor adequate habitat patches for both species; northern

S. griseicollis has been collected as far north as the Ocana

range at La Palmita, Norte de Santander. Other species not

yet recorded in Serranıa de Perija (Lopez-O. et al. 2014)

include S. sanctaemartae and S. latrans, but they occur in

adjacent mountain regions at lower elevations and in

different habitats from those of the new species.

FIGURE 7. Introductory churrs of alternative songs delivered after stimulation with playback: (A) S. perijanus sp. nov., Sabana Rubia,Manaure, Cesar, Colombia, July 6, 2008 (CSA 2818); (B) S. latebricola, Nabusamike, Cesar, Colombia, February 19, 2007 (Krabbe 2008).Advertising songs: (C) S. perijanus sp. nov., type locality, July 6, 2008 (CSA 2822); (D) S. spillmanni, Ucumarı N. P., Risaralda, Colombia,July 17, 2001 (Alvarez et al. 2007).



To better assess the potential distribution of S. perijanus,

and ultimately its conservation status, we conducted an

ecological niche modeling (ENM) analysis in Maxent v3.3

(Phillips et al. 2006) using 19 climate variables (Hijmans et al.

2005), and 13 remote-sensing variables related to vegetation

and 3 related to topography (Buermann et al. 2008).We used

all the localities reported in this paper to build the model

(Appendix C). The distribution model suggests that S.

perijanus is potentially restricted to the northern section of

the Serranıa of Perija between 108510N and 098420N,

apparently tracking the distribution of humid vegetation

above 1,600 m including elfin forests and paramo habitats

(Figure 8). However, the species might be found further

south in the Serranıa de losMotilones. Fieldwork is necessary

along the Motilones in southern Cesar and northern Norte

de Santander in Colombia and southwestern Zulia in

Venezuela to evaluate distributional limits of the new species

and other Scytalopus tapaculos occurring in the region.

Breeding BiologyThe reproductive biology of Scytalopus tapaculos is poorlyknown (Krabbe and Schulenberg 2003, Greeney 2008). We

found and collected a nest of S. perijanus (ICN-N-231) on

July 13, 2008, within a 2–3 m tall secondary forest patch

close to the forest edge, 2 m from the dirt road that crosses

the type locality. Nest location was facilitated by the loud

cricket-like calls elicited by 2 nestlings (ICN 36800, 368001)

and the ejection of fecal sacs. An individual that

approached the nest was an adult male (ICN 36798) with

enlarged testes and no molt, which indicates male

participation in parental care as documented for other

Scytalopus (Greeney et al. 2005, Decker et al. 2007,

Freeman and Greeney 2008, Hosner and Huanca 2008).

Following the terminology of Simon and Pacheco

(2005), the nest was a subterranean cavity with a short

tunnel entrance connected to the chamber, which was

closed and had a globular shape, mainly made of mosses,

grasses, and plant rootlets. Dimensions were 12 cm in

diameter, 9 cm width, and 14.5 cm depth. The entrance

was ~4.2 cm diameter by 10 cm deep and was partially

covered by some stems and leaves of Pteridium aquilinum

(Dennstaedtiaceae), Huperzia sp. (Lycopodiaceae), Sphag-

num sp. (Sphagnaceae), Alloispermum caracasanum

(Asteraceae), Geranium sp. (Geraniaceae), and Lachemilla

sp. (Rosaceae). The area surrounding the nest was humid

with a high density of shrubs and small trees, mainly

Tibouchina sp. (Melastomataceae), Cleome sp. (Cleoma-

ceae), and Achyrocline alata (Asteraceae). There were also

representatives of Baccharis prunifolia, Ageratina cuatre-

casasii (Asteraceae), Relbunium sp. (Rubiaceae), Sphag-

num sp., and Solanum sp. (Solanaceae) among the leaf

litter neighboring the nest.

The nest architecture and clutch size of S. perijanus are

similar to those reported for other species in the genus,

including S. griseicollis (Krabbe and Schulenberg 2003), S.

parkeri (Greeney and Rombough 2005, Greeney 2008), S.

spillmanni (Pulgarın-R. 2007), S. meridanus (Decker et al.

2007), and S. parvirostris (Smith and Londono 2014). Loud

begging calls of Scytalopus nestlings have been reported

for other species (Skutch 1972, Freeman and Greeney

2008) and have also been described as rapid trills or insect-

like in S. latrans (Skutch 1972), S. argentifrons (Young and

Zuchowski 2003), S. micropterus (Greeney and Gelis 2005),

and S. parkeri (Greeney and Rombough 2005).

Our field observations and data taken from specimens

suggest that S. perijanus probably breeds from April to

July. In early May 1942, Carriker collected a laying female,

and several males and females collected from May to June

had enlarged gonads. Fledglings were collected from the

last week of June through July (USNM). This agrees with

our finding of fledglings, a nest, and nestlings in July 2008,

and with reproductive signals shown by most adults

collected at that time (i.e. enlarged gonads, abundant

subcutaneous fat, worn plumage, and slight to moderate

molt). A specimen collected on February 1951 in Cerro

Tamuypejocha on the Venezuelan side is labeled as a

juvenile (PONS 2888; Gines et al. 1953). We did not find

juveniles in February 2009 but registered 3 subadult males

and noted much lower vocal activity than in July 2008.

Ecology and BehaviorLike many other forest tapaculos, S. perijanus is secretive

and difficult to see. The species seems to be common in

dense interior and forest edges of humid and elfin forests

and highland woody bushes in paramo habitat, especially

between 2,500 and 3,000 m elevation. At El Cinco (type

locality), paramo Sabana Rubia and San Antonio, we found

at least 3–4 territorial males in ~2 ha of suitable habitat;

males from at least 2 territories at each site were paired.We observed single individuals foraging in dense thickets

within 1 m from the ground, often using forest edges and

scrubby vegetation along trails (e.g., patches of Rubus sp.).

At Sabana Rubia, individuals often foraged on the ground,

running across grassy open areas between bushes.

Stomach contents from 7 individuals consisted exclusively

of insect remains.

HabitatThe elevational range of S. perijanus covers a variety of

habitats from lower montane to upper montane humid

forest, elfin forest, subparamo, and paramo. Cloud forests

in the Serranıa de Perija, such as those at the type locality,

are typically dominated by Prumnopitys montana, Clusia

multiflora, Ternstroemia meridionalis, Podocarpus oleifo-

lius, Weinmannia pinnata, Illex sessiliflora, and Hesper-

omeles ferruginea (Rangel-Ch. and Arellano-P. 2007).

Annual rainfall averages ~1,230 mm at Manaure, Cesar,

and is seasonally bimodal with slightly drier periods from



FIGURE 8. Map of northern South America showing the location of areas mentioned in the text, point locality records, and potentialdistribution of Scytalopus perijanus sp. nov. (light gray polygons along the Venezuelan–Colombian border). The projecteddistribution of the new species was defined as �0.5 presence probability of the ENM model calculated in MAXENT. Note therestricted range of the new species to the northern sector of the Serranıa de Perija mainly above 1,600 m.



December to April and in July (Arellano-P. et al. 2007).

Vegetation at the treeline inhabited by S. perijanus is

dominated by bushes (Diplostephium floribundum), bam-

boo (Chusquea scandens), and short trees (Hesperomeles

ferruginea and Gynoxys spp.). The subparamo vegetation in

the area surrounding Sabana Rubia is dominated by

bamboo (Chusquea tessellata), grasses (Calamagrostis

intermedia and C. effuse), and bushes (Lourtegia stoecha-

difolia, Arcytophyllum nitidum, Gaylusaccia buxifolia,

Hypericum baccharoides, and H. magdalenicum), with

scattered patches of small trees (Weinmannia pinnata;

Arellano-P. et al. 2007).

ConservationThe forests and paramo of Serranıa de Perija have a long

history of anthropogenic disturbance. Forest cover has

been almost completely cleared on the Colombian slope

from the foothills up to ~2,000 m in La Guajira (Strewe

2004) and especially around the type locality. The remnant

forests above 2,600 m and paramo are highly fragmented

(Rangel-Ch. 2007). The region as a whole has beenhistorically affected by timber extraction and clearing for

pastures, agricultural fields, illicit crops, and aerial

fumigations of the latter (Fjeldsa et al. 2005). Currently,

the main threats to these habitats are cattle raising, fires,

and the abandonment of cultivated land at the tree-line,

which have led to the replacement of upper-montane and

elfin forests by grassland vegetation typical of the paramo,

a process affecting ~37% of the upper Andean forest of the

Serranıa de Perija (Arellano-P. and Rangel-Ch. 2007). No

protected areas exist on the Colombian side of the range,

but ~300,000 ha of humid montane forest and paramo

habitats are protected in the Venezuelan side by the Sierra

de Perija National Park.

The extent to which S. perijanus tolerates anthropo-

genic forest disturbance and landscape fragmentation is

uncertain. Although data on population size or demo-

graphic trends are lacking, we consider this species to be

fairly common in heterogeneous landscapes with primary

and tall secondary forests subject to logging and small

shrubby patches near forest remnants, even in areas

surrounded by exotic plantations (e.g., Eucalyptus glob-

ulosus) and agricultural fields (e.g., coffee). However, we

did not detect it in the deforested and burned slopes that

have turned into tussock grasslands. Although S. perijanus

likely tolerates some level of landscape fragmentation, it

presumably qualifies as Endangered under IUCN criteria

(B1b(i,iii); IUCN 2001), and its small and fragmented range

(,5000 km2) will probably continue declining in extent

and quality.

Implementing effective actions in the Colombian side of

Serranıa de Perija is urgent for the conservation of S.

perijanus and other threatened and little-known birds such

as Metallura iracunda and Asthenes perijana. In addition,

this region is home to other endemic taxa that will likely be

treated as full species in the near future (e.g., Grallaria

rufula saltuensis, Synallaxis unirufa munoztebari, and

Ochthoeca diadema rubellula), or which have been

overlooked (e.g., Myiopagis olallai incognita; Cuervo et

al. 2014). The Serranıa de Perija is recognized by its high

endemism (Hernandez-Camacho et al. 1992, Viloria and

Calchi La C 1993, Lopez-O. et al. 2014), with ~60 avian

taxa largely restricted to montane forests and paramo

habitats and a number awaiting formal description (A. M.

Cuervo personal communication). Likewise, 69 taxa of

vascular plants are endemic to this range (Rivera-Dıaz and

Fernandez-Alonso 2003). Establishing a large binational

protected area, such as the one protecting the Tama massif,

would be possible with the establishment of new national

park or a network of reserves in Colombia connected to

the Sierra de Perija National Park of Venezuela.

ACKNOWLEDGMENTS

We thank our local guides and assistants for their hospitalityand help with field work. For access to specimens, we aregrateful to C. Medina, S. Sierra, and F. Forero (IAvH-A); F. G.Stiles (ICN); J. Dean and C. Ludwig (USNM); M. Lentino, J.Perez-Eman, and J. Miranda (COP); M. Salcedo (PONS); andJ. V. Remsen (LSUMZ). We thank the contributions andsupport of J. L. Perez-Eman (Universidad Central deVenezuela) and D. Lopez and J. D. Palacio (IAvH-BT) forthe genetic sampling, and F. Garcıa for the identification ofbotanical specimens. S. Gonzalez-Caro assisted with the map.We are very grateful to J. Fieldsa for the painting (Figure 4),and to all the recordists that contributed songs to xeno-canto.org. We thank D. Ascanio, M. Lentino, J. Perez-Eman,N. Krabbe, F. G. Stiles, T. S. Schulenberg, and anonymousreviewers for information, literature, or comments thatimproved this manuscript.Funding statement: This study was supported by TheExplorer’s Club, the Facultad de Ciencias at Universidad delos Andes, National Science Foundation DDIG grant (DEB-0910285), the Society of Systematic Biologists, the Lewis andClark Exploration Fund, Grants-in-Aid of Research of theSociety of Integrative and Comparative Biology, the Frank M.Chapman Memorial Fund of the American Museum ofNatural History, the Alexander Wetmore Memorial ResearchFund of the American Ornithologists’ Union, and the LouisAgassiz Fuertes Award of the Wilson Ornithological Society,and IdeaWild. The first trip to Perija by A. Cortes-Diago andJ. P. Lopez-O. was supported by Ministerio del MedioAmbiente, Corpocesar, Corpoguajira, Conservacion Interna-cional-Colombia, and Fundacion EcoHabitats. The visitorfellowship program of the Smithsonian Institution supportedJ. E. Avendano’s visit to USNM. None of our funders had anyinfluence on the content of the submitted or publishedmanuscript, neither require approval of the final manuscriptto be published.Ethics statement: Collecting permits were obtained fromCorpocesar (Resolucion 0961).



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APPENDIX A

Specimens of various taxa in the genus Scytalopus

examined at the Coleccion Ornitologica Phelps (COP),

Coleccion Adolfo Pons (PONS), Coleccion de Aves del

Instituto Alexander von Humboldt (IAvH-A), Instituto de

Ciencias Naturales (ICN), the National Museum of Natural

History, Smithsonian Institution (USNM), and LouisianaState University, Museum of Natural Science (LSUMZ).

Adult and juvenile specimens are included. Specimens

included in phylogenetic analysis are highlighted with an

asterisk.

S. latebricola (all from Magdalena, Colombia). Males

(17): IAvH-A 2049, ICN 36194, 36257*, USNM 170554,

387414-5, 387417, 387420, 387422-25, 387429-31, 387433,

LSUMZ 90420; females (14): ICN 23338, USNM 170553,387412-4, 387416, 387418-19, 387421, 387426-8, 387432,

387434.

S. perijanus sp nov. Males (25, not including the type

series): USNM 369262-3, 369265, 369267, 373421,

373423, 373425, 373430, 373431-2, 373437, 373438 (La

Guajira, Colombia), COP 54931, 54938-40, 54945, 72846-

7, 72851-2, 72854, 74172, PONS 2890-1 (Zulia, Venezu-

ela); females (20): USNM 369261, 369264, 369266,369268, 373420, 373422, 373424, 373426-7, 373429,

373433-36 (La Guajira, Colombia), COP 54933, 54947,

57708, PONS 2889, 2892-3 (Zulia, Venezuela); unsexed

(3): ICN 36045, 36046 (Cesar, Colombia), USNM 373428

(La Guajira, Colombia).

S. meridanus. Males (16): all from Venezuela: COP

14363, 14577, 45379, 45382, 49294-5, 73947 (Merida),

9441, 11102, 24551, 83654-5, 84104, 84105*, 84108*, 84109

(Tachira); females (4): all from Venezuela: COP 14205,65397, 71525 (Merida), 84107* (Tachira); unsexed (2):

COP 84106, 84110 (Tachira).

S. caracae. Males (17): all from Venezuela: COP 13051,

13153, 13155, 56785, 58461, 58466, 58468, 58471-72,

61643, 61645, 62230, 62610, 62612-3, 62619, LSUMZ

14194 (Caracas D.F.); females (10) all fromVenezuela: COP

13049-50, 56782, 58464, 61648, 62606, 62609, 62616,

62618 (Caracas D.C.), 75775 (Aragua).S. griseicollis. Males (19) all from northeastern

Colombia: IAvH-A 8416, 14919*, 14841, 14946, 14797,

14948, 12068, 12125*, 10728, 10664, USNM 373414

(Norte de Santander), ICN 36121, 36416, 37514, 37538,

37548, 37570, USNM 402075, 411789 (Santander);

females (12) all from Colombia: IAvH-A 8427, 8434,

8438, 10625, 12123, 14941, USNM 373415 (Norte de

Santander), ICN 37516, 37522, USNM 313714, 402074,

411790 (Santander).

APPENDIX B

Recordings of primary songs and calls of Scytalopus

perijanus sp. nov., S. latebricola, S. meridanus, and

northern S. griseicollis used in quantitative analyses of

vocalizations. We used the average of 2 to 4 song bouts

per individual for S. perijanus and northern S. griseicollis,

and the values from one song per individual for S.

latebricola and S. meridanus. Two calls per individual

were analyzed for S. perijanus, whereas 1 to 2 calls per

individual were considered for the other species. Alter-

native and advertising songs of S. perijanus are also listed

below. Acronyms for sound archives: CSA, Coleccion de

Sonidos Animales-Instituto Alexander von Humboldt,

XC, xeno-canto.

Primary SongsS. perijanus: Colombia, Cesar, Manaure, Sabana Rubia (J.

E. Avendano, CSA 2801-2; J. P. Lopez, CSA 2803; A. M.

Cuervo, CSA 2804-5); Colombia, Cesar, Manaure, San

Antonio (A. M. Cuervo, CSA 2806-8). S. latebricola:

Colombia, Magdalena, Santa Marta, San Lorenzo ridge

(Krabbe 2008, # 1, 2, 4, 7; Strewe et al. 2005, # 27; C. Hesse,

XC10186); S. meridanus: Venezuela, Tachira, Paramo del

Zumbador (Boesman 1999, # 2.1); Venezuela, Merida, 10

km SE La Azulita (C. Parrish, XC6236; A. Spencer,

XC14790); Venezuela, Merida, Sierra Nevada N. P., Pico

Humboldt trail (D. Edwards, XC20071); Venezuela,

Merida, Sierra Nevada N. P., La Mucuy (J. Klaiber,

XC43158, XC43160); S. griseicollis (northern): Venezuela,

Apure, Rio Oira (C. Parrish, XC16656); Colombia,

Santander, California, Angosturas (J. E. Avendano,

XC86713-15).

CallsS. perijanus: Colombia, Manaure, above El Cinco (A. M.

Cuervo, CSA 2809; J. E. Avendano 2810); Colombia,

Manaure, Sabana Rubia, Casa de Vidrio (J. E. Avendano,

CSA 2811-13; A. M. Cuervo, CSA 2814-15); Colombia,

Manaure, San Antonio (A. M. Cuervo, CSA 2816); S.

latebricola: Colombia, Magdalena, Santa Marta, San

Lorenzo ridge (Krabbe 2008, # 9, 11; C. Hesse, XC10187;

N. Athanas, XC10746; D. Geale, XC51259); Colombia,

Cesar, 3–8 km NNW Nabusimake (Krabbe 2008, # 10, 19);



S. meridanus: Venezuela, Tachira, Paramo de Batallon

(Boesman 1999, # 2.4); Venezuela, Merida, Sierra Nevada N.

P., Loma Redonda-La Aguada (B. Lopez-L, XC50485-86);

Venezuela, Merida, Sierra Nevada N. P., La Mucuy (B.

Lopez-L, XC50641); S. griseicollis (northern): Venezuela,

Apure, Rio Oira (C. Parrish, XC6079, XC16657, XC16659,

XC16661); Venezuela, Tachira, Betania, Tama N. P. (A.

Renaudier, XC23330-32); Colombia, Santander, Guasca,

vereda Concepcion (O. Cortes, XC27187); Colombia,

Santander, Surata, vereda Bucare (J. E. Avendano,

XC86716-17).

Alternative songs.—S. perijanus: Colombia, Manaure,

Sabana Rubia, Casa de Vidrio (J. E. Avendano, CSA 2817-

18; A. M. Cuervo, CSA 21819); Colombia, Manaure, El

Cinco, above El Cinco (A. M. Cuervo, CSA 2820-21).

Advertising songS. perijanus: Colombia, Manaure, above El Cinco (J. E.

Avendano, CSA 2822).

APPENDIX C

List of localities of Scytalopus perijanus sp. nov. (geo-

graphical coordinates in decimal degrees and elevation

above sea level) used in ecological niche-modeling analysis

(Figure 8). Locality data were obtained from museum

specimens, song recordings and reliable observations.

Colombia, La Guajira: Sierra Negra, Monte Elıas (10.832,�72.686), 1,530-1,830 m; Tierra Nueva (10.614, �72.801),1,370–1,830 m. La Africa (10.524,�72.935), 1,615 m. Cerro

Pintado, Laguna de Junco (10.445, �72.902), 2,290–2,740m; Cerro Pintado (10.5, �72.887), 3,230 m; Cesar:

Municipality of Manaure, Sabana Rubia, Casa de Vidrio

(10.367, �72.897), 3,030 m; El Cinco, above El Cinco

(10.364, �72.953), 2,450 m; San Antonio, Finca Villaluz

(10.364, �72.993), 1,600–1,800 m. Camp above Hiroca, SCerro La Teta (9.93, �73.046), 1,675–1,980 m. Cesar:

Municipality of La Paz, corregimiento de San Jose de

Oriente, Brisas del Perija (10.262,�72.962), 2,800–3,000m.

Venezuela, Zulia: Cerro Viruela, NW foot Cerro Pintado

(10.457, �72.884), 3,180–3,230 m; Campamento Frontera

5, foot Cerro Pintado (10.439,�72.862), 2,710 m. Serranıa

de Las Lajas, Antenas (10.437, �72.738), 1,800-1,850 m.

Campamento Avispa, Fila Macoita y Apon (10.403,�72.837), ~2,000 m. Pie Nudo 4 (10.394, �72.792),~1,900 m. Cerro Tamuypejocha or Pejochaina, Cumbre y

falda (10.095, �72.815), 2,300 m. Rıo Negro, Manchique

abajo (10.071, �72.847). Cerro Tetare or La Teta, Pie SE

(10.036, �72.935), ~2,900 m. Campamento Frontera 2

(10.029, �72.878), 2,270 m.



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